HOW  DO  THE  MODIFIERS  OF  LINKAGE  VALUES  AFFECT 
DETACHED  PORTIONS  OF  THE  CHROMOSOME? 


BY 


WELLINGTON  SHANG  HSU 


THESIS 


FOR  THE 

DEGREE  OF  BACHELOR  OF  SCIENCE 


IN 


AGRICULTURE 


COLLEGE  OF  AGRICULTURE 
UNIVERSITY  OF  ILLINOIS 


1922 


Digitized  by  the  Internet  Archive 
in  2015 


https://archive.org/details/howdomodifiersofOOhsuw 


UNIVERSITY  OF  ILLINOIS 


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CONTENTS 


PAGE 

I IJaTRODUCTION 1 

II  MATERIALS  AITO  METHODS 4 

III  THB  DATA  AND  DISCUSSION 

A-  REGION  l<5b2 8 

B-  REGION  4 13 

IV  CONCLUSIONS 13 

V ACKNO’IPLEDGIMENT  17 

VI  literature  cited 18 

VII  TABLES 

A-  TABLE  I 3 

B-  TABLE  II  9 

C-  TABLE  III  AND  IV 14 

D - TABLE  V 19 

E-  TABLE  VI  21 


F-  TABLE  VII 


23 


iV 


I 


HOW  DO  THE  MODIFIERS  OF  LIHKAGE  VALUES  AFFECT  DETACHED  PORTIONS 


OF  THE  CHROMOSOME? 

I 

nJTBODUCTION 

The  axnooint  of  llteratixre  that  has  been  published  recently  bearing  on 
the  phenomenon  of  linkage  of  hereditary  factors  makes  it  unnecessary  to  give  any 
description  of  the  same  at  tills  time.  To  eiQ)lain  the  vest  accumulation  of  data 
obtained  from  intelligently  planned,  and  carefully  controlled  experiments,  several 
theories  have  been  advanced;  but  as  time  goes  on  it  is  becoming  more  and  more 
obvious  that  the  chiasmatype  theory  is  the  most  plausible.  According  to  the 
chiasmtype  theory,  Mendelian  factors,  or  genes,  reside  in  a linear  series  on  the 
chromosomes,  and  each  has  a definite  locus  with  its  allelomorph  occupying  the 
corresponding  locus  on  the  other  member  of  the  homologous  pair  of  chromosomes. 

The  immediate  cause  of  crossing  over  is  attributed  to  the  breaking  of  the 
homologous  pair  of  chromosomes,  which  twist  arour.d  each  other  during  synapsis, 
and  the  subsequent  reunion  of  them,  each  taking  a piece  that  formerly  belonged  to 
the  other. 

Taking  this  theory  as  a working  basis,  geneticists  had  accismulated 
sufficient  data  to  convince  themselves,  until  quite  recently,  that  the  crossover 
value  between  two  Mendelian  factors,  is  constant  and  invariable*  It  is  the 
amount  of  such  data  and  the  accepted  concept  of  the  linear  arrangement  of  genes 
on  the  chromosome  that  led  Morgan  to  believe  that  crossover  valtie  between  two 
linked  factors  is  a fhncticn  of  distance.  Attractive  and  suggestive  as  Morgan's 
postulate  may  be,  it  has  not  been  wholly  successfhl,  how'ever,  in  piecing  itself 
beyond  question.  New  searches  were  conducted  by  different  investigators  and 


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th«y  have  showi  that  cxx»saover  values  are  variable,  being  Influenced  by  age 
(Bridges  *16),  t^s^ierature  (Plou^  *17),  and  also  certain  specific  genetic  factors 
(Sturtevant  *17), 

The  array  of  evidence  against  Morgan's  view  was  amplified  by  the  work 
of  Detlefsen  and  Boberts  (Datlefsen  and  Hoberts  *21)  tnho  worked  on  Drosophila, 
and  through  selection  have  been  able  to  reduce  a crossover  value  of  32^  (from 
white  to  miniature),  to  practically  zero,  in  one  of  tl^e  series  at  the  end  of  the 
12th  generation.  Instead  of  accepting  Morgan's  hypothesis  of  distance,  Detlefsen 
C2I)  propounds  that  the  linkage  value  is  at  least  to  a large  extent  "determined 
by  the  different  possible  combinations  of  uultiple  modifying  factors."  In  order  to 
test  out  this  view,  he  actually  performed  a hybridization  experiment  between  the 
low  crossover  stock  and  normal  flies,  A portion  of  his  data  is  reproduced  in 
Table  I.  In  the  table,  there  ere  to  be  found  in  the  first  and  second  lines  the 
distributions  of  crossover  values  of  normal  populations  and  generation  of 
series  B,  In  ^42  generation  of  series  B low  stock,  he  mated  fifty  red  long  females 
to  normal  white  miniature  males,  and  the  crossover  val'ues  of  these  females  are 
shown  in  the  third  line.  To  secure  a representative  distribution  of  crossover 
values  of  the  Fp  hybrid  females  thus  derived,  he  chose  four  such  females  fTom 
each  of  the  forty- four  pairs  and  crossed  them  to  their  brothers.  The 
distribution  of  such  a population  is  given  in  the  fourth  line.  It  is  stated  that 
since  it  is  impossible  to  test  out  the  crossover  capacity  of  the  male  parents,  we 
mist  accept  for  them  the  value  of  series  B Fp  generation,  since  they  came  from  a 
stock  which  had  been  used  for  class  work  for  many  years  and  always  found  to  give 
a value  approximating  33^,  The  Fg  distribution  shown  here  is  the  total  of  three 
distinct  and  separate  F2  distributions  which  came  from  P^^  pairs  No.  2,5  and  6. 

It  is  almost  needless  to  point  out  that  the  F^  distribution  lies  in  between  those 
of  the  two  parents  and  so  do  both  the  value  of  the  mean  variate  and  mean  total 


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crossover  value.  The  T2  has  a wider  range  of  distribution  than  that  of  the 
but  the  values  of  the  mean  variates  and  the  mean  total  crossover  values  is  about 
the  same  as  the  Fx,  Such  features  characterising  a hybridization  eij>eriment  vdiere 
multiple  factors  with  incoB^lete  dominance  are  involved  are  further  confirmed  by 
the  increased  standard  deviation  of  the  Fg  distribution.  It  is  this  and  other 
genetic  analyses  that  led  Detlef sen  ( *21)  to  wonder  "Just  sdiat  part  distance 
between  two  genes  on  a chromosome  map  may  play  in  determining  linkage  values.  " 

The  present  work,  vdiich  is  really  a minor  part  of  his,  was  ^undertaken 
with  the  hope  of  throwing  further  light  on  the  whole  problem  by  attempting  to 
find  out  how  the  modifiers  of  linkage  values  affect  detached  portions  of  the 
chromosome.  Would  the  separated  pieces  still  give  low  values,  or  would  they  be- 
have in  some  other  fashion? 

II 

MATERIimS  mB  METHODS. 

To  cope  with  the  coiqplexity  of  such  an  experiment,  Professor  Detlef  sen 
developed  a special  system  of  designation  Which  I shall  use  in  the  following 
description.  According  to  this  system,  the  piece  of  chromosome  between  vdaite  and 
miniature  is  divided  into  three  different  regions  as  shown  In  Figure  1.  Individu- 
als carrying  the  different  pieces  of  chromosomes  are  named  accordingly.  For 
instance,  individuals  carrying  the  piece  of  chromosome  from  red  to  cross-veined 
end  the  rest  of  the  chromosome,  comes  from  the  other  member  of  the  homologous 
pair  is  considered  as  Region  1.  Those  carrying  the  pieces  from  cross-veined  to 
not-cut  is  called  Region  2,  etc.  The  shaded  portion  in  the  figure  represents 
the  region  outside  of  selection.  With  reference  to  this  region  a more  detailed 
llscusslon  will  appear  in  the  latter  part  of  this  paper. 


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involved  in  the  experiment,  and  Uia  meaning  of  "Region".  The  shaded 
portion  represents  the  region  that  is  outside  of  selection  influence. 


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The  origin  of  our  experiiTiental  materials  was  as  follows:  Mr.  L.  S, 

CleiQentd  crossed  normal  homozygous  recessive  females  of  the  zygotic  constitution, 
WjCjChiinif fdiite  (eye)  cross  veinless  (wing),  cut  (wing),  and  forked  (bristle) 
to  a dominant  male  of  the  genetic  formula,  W,C,C'b)^*^  (red,  cross-veined,  not  cut, 
long,  and  not  forked)  which  he  obtained  from  Prof.  Detlefsen’s  low  crossover 
stock  F7g,  As  usual  he  secured  homozygous  recessive  males  and  dominant  females 
heterozygous  for  the  factors  involved:- 

wcchmf  9 wccbmf  <f 

Mating  the  sibs  together,  crossovers  in  vario\u»  regions  weze 
obtained,  Mr.  Feldman  took  over  the  females  of  Begion  1 (ib  2,  and  Region  4 was  run 
yss  myself.  Among  these  various  region  crossovers,  we  secured  those  of  Begion  1 2 

and  those  of  Begion  4.  The  process  in  which  they  are  supposed  to  have  evolved 
is  shown  in  the  following  diagram;- 

Begion  1 (!b  2 

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w c ch  m f 

W C m f 


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A list  of  the  classes  of  offspring  obtained 
and  Heglon  4 females. 


Parental  — 


Crossover  — — 


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Double  Crossovers 


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-is- 

Theee  eroseovers  made  up  the  material  of  Experiment  1. 

Experiment  2 had  a somvvdiet  elmllar  origin.  In  Experiment  2,  the  male 
that  originated  the  population  was  from  Eg^  generation  of  Dr,  Detlefsen's  low 
crossover  stock  and  the  females  to  which  it  was  mated  came  from  a quadruple 
recessive  stock  v^lch  had  been  subjected  to  six  generations  of  inbreeding.  This 
quadruple  recessive  stock  was  a white,  cut,  cross  veinless,  miniature. 

Our  methods  of  procedure  with  the  Eg  segregates  of  the  various  regions 
are  the  same, for  females  carrying  any  given  region  were  mated  to  the  ultimate 
recessive  males.  The  different  classes  of  offspring  resulting  from  these  two 
crosses  are  shown  in  the  accompanying  chart.  Each  female  was  kept  with  three 
or  four  males  of  the  right  kind  in  8-drachm  homeopathic  vials.  The  common  method 
of  culture  for  Drosophila  was  followed.  Elies  were  r^sooved  to  new  vials  every 
other  day  and  a new  supply  of  males  was  introduced  to  take  the  place  of  the  dead 
ones,  should  deaths  occur.  Nine  days,  from  the  day  when  the  first  offspring 
emerged,  were  allowed  for  the  last  one  to  be  hatched  out  in  each  vial,  and  three 
countings  were  made  during  the  nine  days.  All  calculations  were  done  twice  on 
one  of  the  best  calculating  machines  on  the  market,  end  it  is  hoped  that 
arithmetical  errors  have  been  reduced  to  a minimum. 

Ill 

DATA  AND  DISCUSSION. 

A - Region  1 and  2. 

The  distributions  of  cross  over  values  for  Region  1 and  2 in  each 
experiment  separately  and  combined  into  a total  are  given.  They  are  plotted  in 
class  intervals  of  1.5^.  The  distribution  for  Experiment  1 ranges  from  0 to  18.5^ 
and  for  Experiment  2 from  0 to  14.25^.  The  total  of  the  two  combined  is  shown 
in  the  last  line  in  the  table,  and  the  curves  plotted  from  them  are  shown  in 


t ‘ 

<N^.y  f '( 

t •:  . . 

« ' I ■»  'i.  ' 

J ■' 

•V,  t' 

I - ‘i 

.’.a, 

.::  l!’ 

t . . 

< LO  -h  i.  - 


t ii:' 

i.\ 

dfc  4? 

a I 


’.-wwp 


t -i  TJrf  i-| 


“'  'if. 


4i*  '*  v^.  ~ " f » 1 

• .vieVt 

..^  '"'  ' ' , J ' * 


i ^ 
!■ 


■ ■■  '-  ^,.  •'  ■; 

f V !-_?■  “•:  ■*:  » '.'  I'.w  > x 5J^ 


A A- 


r^r 


v:  r 


m 


ii  »'•*■  til'-  (,-«:»*. 

;.  ^ w *-•  - ■ II-,  . J ' i ■ 


,«  ^ 


//•‘i  -j 


W% 


1^.. 


fr, 


^ * 


;•»,  • 'r'  ».'  >Y 

',  -I , „Y  . . •;.  '■  .»■•«  j»-‘*..3rv  ^ ' •;.'  l>\  'Tfi  ,p|[^i%'  s-|;'"'a||W 


“■;.L  '.>  ^Sr  V 

,»x.«  L '•  *.♦  > 


lUd.  15!>‘»9  •>: ; «ikttf  f :.»*i'ltl<>:  4k'  ». 

* t ji  ' 


,‘r 


t/i  ♦ i*' 


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.’v  , ' _'®'.  •** 


A. 


I 


' . ‘ ■ ':■£  .*  .IL  ,-rL-  ,1 


a '■Xf™') 
£ •'1; 


£ 


-'•  -s-  .■-••  " - -5  ..  ■ ' ^ ^ 

•.,.r,<-r;  ,J4J,,  .-Vt  Kift  -j;? 

..,  ipV.  . ' * 

' f^7; 

••t-'ji--;*  4'  *f  t T 

1/ 

iVl  ' , ® *f 


: ( ' ■■  h_4  * -'_- 


• . 

• '/  «V>  V 


, L ^ » *7 


f ^ V 


. k 

Iji 


' w ;• 


jc-:  -.  i 


s.  4 ‘i^.«7i'**  ' * xCii  V ••  **’*  - 

i ;:.  iv-'i  tis.;  * ’•■  ■ 

- --  .,.n .,,  j.3*j4Hi»ji 


^ U'  •••  ‘ ; 


. (O'!*.  i/W  sf 


r »i;j^  - ^ - 


; .{ r 


ifU 

r-=*sfc  .rrrtuT'^^amsrStss*..^^ 


TABLE  2 


CM 

«« 


a 

O 

•ri 

O 

PEI 


% 

® CD  (Sl 

s > 


lol 


lO 

CM 


lO 


(0  m 

N. 


a 

> 

s 

>• 

o 

a 

« 

o 

»4 

o 

«H 

o 

a 

o 

ft 

*> 

•H 

«4 

4> 

.9 

« 


*M  • 

o ® 


55  ® 


$3 

! o 
j*^ 
!■** 


o 

o 


(3 

o 

•H 

4* 

d 

>4 

O 

a 

o 

C9 


Si 


^ 9 
P ^ 

«H  +> 


P 

V 

•H 


d 

o 


t4 

► 
o 
« 

o 

d 

o t* 
9 

I ® 

^ q> 

FI  to 


O r-« 

-»>  o 

rO 


U 
9 
> 

CM  O 

e 

• « 

to 
® 

P3  u 


o I 
> 

•H  O 

9 

• a 

to 

© >« 

« o 


to 

»<  © 

0<  © rH 

i “•! 

Vi  © 
O V» 

© 

> 

o © 

« n 

« (H 

o © 

► 

o 


« 

h 

© 

► 

O 

• 

tt 

o 

M 

O 


*3 

4» 

o 

E4 


o» 

O 

• 

00 


'I* 

o 

CO 


vO 

CM 

a* 

CM 

e- 

« 

• 

• 

CM 

CM 

CM 

00 

tn 

• 

• 

• 

CO 

CO 

CO 

in 


g g 


CO  00  r-l  e-  O 
m 00  CO  in  o» 

vo  LO  vo  m U3  <x>  cn 


o fH  CM  CM 
CM  CM  e-  CO 
CO  O O CM 


g 


CO 

vO  C-  CM  CO 
00  MS  M3 
in  o 00  CO 


g 


CM 

in 

in 


g 

<j» 


O to  ^ 00  ^ 


to  <»•  O'  00 


8 


M to  «0 

SO  O' 


0 

1 


o 

o 

IP 

CP 

o 

< 

Q> 

►1 

•4 

(0 

IP 


Experiment  II 
Experiment  I ft  II 


-11- 

Figare  2,  The  most  striking  feature  of  the  curves  is  the  large  numher  of 
individuals  in  the  first  class,  making  them  look  rather  unique  for  the  experi- 
ment. However,  if  we  turn  to  the  hack  and  look  thrcu^  Tables  5 and  6,  we  will 
find  that  there  are  quite  a few  of  the  F£  segregates  giving  such  a small  number 
of  offspring  that  their  crossover  values  reached  zero.  It  is  true  that  zero 
percent  crossover  value  should  be  expected  to  occur  in  this  case,  if  we  ejq)ect 
modifiers  of  lixkage  values  to  segregate  at  all;  but  still  we  should  not  over- 
look the  fact  that  over  half  of  the  individuals  that  give  zero  percent  are  those 
that  have  exceptionally  small  number  of  offspring.  Therefore  due  allowance  must 
be  made  on  the  first  class  in  Table  2,  and  thus  the  medal  classes  of  the  curves 
could  be  shifted  to  5.25^  for  Es^eriment  2 and  3.75^  for  E:q>eriment  1 and 
Experiment  1 and  2 combined. 

From  the  description  given  in  the  early  part  of  this  paper,  it  is 
very  obvious  that  we  have  also  performed  a hybridization  es^eriment  on  two  strain: 
of  Drosophila  differing  from  each  other  in  their  capacity  for  crossing  over.  In 
fact,  as  has  been  shown,  this  has  been  admirably  done  by  Prof.  Detlefsen  him> 
self.  The  only  point  in  which  his  work  differs  from  mine  lies  in  the  fact  that 
he  \ised  the  Tdiole  region  between  white  and  miniature;  while  I dealt  with  se^ents 
of  it.  Due  to  lack  of  experience  in  handling  the  material,  and  in  the  method  of 
culture,  the  results  obtained  could  be  much  better.  There  are,  however, 
reflected  in  the  data  a few  features  that  are  characteristic  of  a hybridizing 
experiment  vhere  multiple  factor  Inheritance  with  Incomplete  dominance  is 
involved.  These  features  stand  out  so  strikingly  that  it  is  but  Justifiable  to 
arrange  the  material  and  present  it  in  its  logical  shape.  Owing  to  the  fact  that 
Prof.  Detlefsen  is  dealing  with  the  whole  piece  of  chromosome,  it  is  impossible 
for  me  to  find  out  the  crossover  values  of  Beglon  1A2  for  the  respective 
generations  to  which  the  two  originel  males  belong.  However,  we  know  the  value 
for  the  whole  region,  and  in  order  to  present  the  data  in  the  shape  it  is  now. 


h-AMM 


t. 


^ J.*?  - - i 

B..  ' ' . . ^ -.  '^"  ■■ 

i-W:  ^i‘  I'iitti*.  f'.  .V*>.v'.'  ':-Hi’t»i 

'i  *!#tv  -<?■  ;aW-  :■•’••*•  •»'• 

% "|gc 


"t 


f i , -i'  . * 

'■  ;";^#Pr 

.;-ii't..  ■■mi  „ nr  i 


,.  * r i •* ,.: 


I ''it  , ' . '••  JT*. 

^1  J^*‘  ' , . ^ ■•  " ^ t U JBHV  1 1 


*>•’  ' 1^  ■' |,  . ' ' Ja ' ••  >■'"*“  ' 

■’  ; V’ " ^ ^ • : vt,‘/|iP'  ip(^',  >r^v  Uf'  ■ ‘ 


^ ,'  ■■''5  ft’tv  y:  ''A  ■■-■•-  • 'V  ■''/*•'•■' 

.*-.!’•/  ■ / »-li  t «•  •• 

, .'  - .;vj  risiiC  i ' ■ S - 


»V' ,:  '.'■■xii' 


•/»'®  1 


li!  ' ■ •*•' 

t. 


0*/  ■-r-v'i’  L 

' .'  ' * ^ -4  ■'■''-•  4'f  ■..  •■•"'■•  i'*^'  r 

•::  , . ■>•  ■:■.  J --5 

.. S .. J-  ! ,t  ■ ■ •• . 


!ts 


? ' ''  • 

-w  ■f  . i.*r»  . ,.< 


iiia‘  ■'■*  % T|i,»«-' 


. 'r’’\^V  '''''  .'^  v'?^LW,  -Bix-r- 


d--^::.  i?.: 


■.  " ■^■T<^“  ^ • ■ ,.j 

.,ij  !y  , ‘<i^,J'.i.^s.  „.»,■:, -V,  .,!,■,  .M/4.»i 

*’^''  ,i'  d'O'  '^,'-1  .* 

■ i>  I'  r -V  ' * '*‘  '**'*•:  ■^  *•,  ■<■“;■  •'  |^,*>  * «■  '.«*>  • f/  "■' . ■ , f jj^ 


yt'13 


i'' 


n 


r i.  .■  i'd  ,;c, -4^<. 'fe 


.3..-..-..  i .sx*.  f 

. .!l  i'.«  ,i  ■'  ,M^ri  4"  *V<J».t‘-')iy 


i'  “ ,^.-.iv , .Tj  i.  .T.-V-.  -T.  • . v.^  .-  , ^ 


, '■  ’•  ■'  ■ . ■''-....  ' 
lOffin-.'"  .-'ll'  ■ “‘■i-Xt'J.tf'XXt'’'*"” 

■ - - 1 -4^  • - 1^  t r ^ ® 


7<r 


,,  ,.  ’.  , ; . ,:j,  «J  »»?« 

. f .-jd' 

0? 


ir  . . u'.»j  iaoin'ii  .•.»  ‘ 


if.* 


t. 


f : ■'  \ . 


,:  , Vv  --‘ft*: S4iXi*« ••3 

..-i  ^ ,..,  . • :<  lit  ZU  atiy^\  '<• 

■•«  c - ' V-..  lE'  . 

^ ..d  ,7l|  ■'#  ' ,V, ;■, 

i‘- ,rr9^*'  r;‘‘rrn-^'^.ri^'iSi‘j:,twa*r'  <^SSS!Xl^^ 


r 

•12- 

It  is  Isperatlve  that  we  assign  to  Beglon  1 (ib  2 a yalua  which  we  think  most 
nearly  true.  This  value  is  indicated  hy  a cross  in  the  first  line  of  Table  2. 

The  females  used  were  normal,  being  considered  capable  of  giving  18.5^  of  cross- 
overs within  Beglon  1 cSb  2.  From  a cross  of  this  description,  we  naturally  e:^ect 
an  Fx  population  giving  an  intermediate  value  of  crossover  between  the  values  of 
the  parents.  This  opportunity  for  testing  the  F^  generation,  however,  was  not 
given  to  the  writer  but  throu^  the  kindness  of  Mr,  Clemente,  the  F^  crossover 
values  for  both  Experiments  were  obtained  which  fare  shown  in  the  table. 

It  is  readily  seen  that  the  F^  crossover  values  of  both  experiments  are 
not  hi enou^  to  be  called  intermediate  between  the  parents.  Those  of  the  F£ 
are  still  lower,  A clearer  idea  of  the  data  is  better  grasped  by  glancing 
over  the  curves  in  Figure  2.  If  due  allowance  were  made  in  the  first  claases,  we 
can  but  conclude  that  a partial  dominance  of  the  ”low  modifiers"  must  be  existing, 
since  the  mean  total  crossover  value  of  the  F^  and  Fg  is  lower  than  an  arithmetical^ 
mean  between  the  original  parents. 

Now,  if  we  compare  the  F£  crossover  values  of  detached  piece  with  those 
of  v^nole  chromosome,  we  will  find  that  they  show  up  in  the  same  general  magnitude, 
being  5.44^  in  Experiment  1 and  15,37;^  in  Experiment  2 as  compared  to  5,817^  when 
the  vdxole  chromsoma  was  involved.  This  direct  comparison  of  Fj  and  F2  data  brings 

out  clearly  the  fact  that  modifiers  of  linkage  values  affect  detached  portions  of 
chromosome  same  as  they  do  the  vdiole  piece. 

Since,  as  has  been  mentioned,  the  low  male  used  in  Experiment  2 was 
from  F34  generation  and  the  females  0B5>loyed  had  been  subjected  to  six  generations 
of  inbreeding,  vhile  those  used  in  Experiment  1 were  just  nonnal  individuals  and 
the  male  was  from  Fyg^  ^3  8ho^lld  expect  the  standard  deviations  of  the  two  F 

2 

posalatlon,  to  Show  soma  diffaranoa.  A glanoa  at  that  oolwm  In  the  table  will 


'4 


■ ' ■/"  ■■•  ;-v  ' sft 


'M'  '.  ^ '' 


. 

4.;^ 


r 


y , 


t f-i  ■ - v'V‘.\i. 


Vm.  ; , , 

■•  , J ;v  . ■<  V ■- 

, . •!!' 

j, . ,,  • ' i l ■*■  . I,  ' 


- - :;f  ' • ' 


r .'■  . -'V  w- 

."■j't  'J.J  ■ J ■ ■'“  - 1 •• 

I ■:  "I  ,.v  '■■  ■!  t 

. •.•  ■ A 

, r t-i'f •V'-  i-H- 

C'  t',C  ..i'  “•  *'•  • * 

' - r * . 

' ■^  7 ’ • •■  < 

• i’-  " ■ . ' ' 

. i.  ■ ■ ■"'rf  ■ •. 

, t..,  ~ '■■ 

I t.  .. 

;.  J I ■';  V(t  >1 

••\(i.  .>  1 '■  '*  ’■’  ■ ' ''  ‘ 

' ■ 


•t^  -ipt  -“'f^ 


13- 


tell  that  they  do  differ,  and  also  in  such  a wai'  as  we  eaqpected.  The  standard 
deviation  of  Experiment  1 being  bigger  than  that  of  Ej^)0riment  2. 

B - Eegion  4. 

As  mentioned  at  the  beginning  of  this  paper  the  purpose  of  this  work 
is  more  of  the  nature  to  try  to  throw  farther  light  on  the  whole  problem  of 
modifiers  of  linkage  values  than  merely  to  secure  more  data  to  expand  the  theme 
stated  in  the  subject,  so  I ran  a test  on  Region  4 vdxich  is  the  space  between 
miniature  and  forked,  and  it  is  outside  the  original  region  of  selection.  As  it 
appears  in  Table  3,  this  region  does  not  seem  to  have  been  influenced  by 
selection.  Forty-one  faaales  were  tested  and  with  an  average  of  127  offspring 
per  female,  the  crossover  value  obtained  for  the  region  is  19,75^,  vdiich  is  very 
close  to  the  recognized  valuet-  20.4^.  Owing  to  the  extremely  small  number  of 
offspring  obtained  from  some  of  the  females,  the  distribution  table  drawn  up  is 
rather  puzzling.  Since  selection  does  not  seem  to  have  any  effect  on  this 
region,  the  Fg  population  should  show  a normal  curve  varying  arounii  the  recogniz- 
ed value.  Too  much  emphasis, however,  shoxild  not  be  laid  on  the  distribution  we 
obtained,  since  gi^ite  a number  of  the  females  give  so  few  offspring  that  the 
degree  of  reliability  of  their  crossover  capacity  as  reflected  in  the  data  is 
doubtless  deceiving.  To  eliminate  this  difficulty,  females  giving  less  than 
fifty  offspring  are  left  out  of  consideration,  and  the  resulting  distribution 
is  shown  in  Table  4,  A curve  plotted  from  this  distribution  is  shown  in  Figure  3 
with  17J^  as  the j modal  class. 

IV 

CONCLUSIONS 

From  the  above  discussions  of  data  the  following  conclusions  may  be 


drawn. 


Plotted  from  Table  IV 


Individual 


O ' to  1/1  O' 


o 

at 

3 

ef 

o 

o 

<0 

« 

o 

< 

(b 

m 


Figure  3. 


-16- 


1  Segnant#  of  chroaooome  when  separated  from  the  vihola  piece 

of  chromosome  under  selection  still  show  low  crossover  value  - a fact  showing  t’nat 
modifiers  of  linkage  values  affect  detached  portions  of  chromosome  in  the  seme 
fashion  as  they  do  the  vriiola  piece. 

2  In  a hybridization  experiment  performed  on  "low"  and  "normal" 

flies  with  only  a segment  of  the  idiole  piece  of  chromosome  under  selection,  the 
mean  value  of  the  generation  did  not  come  out  quite  as  hi^  as  would  he 
expected  on  the  basis  of  incooplete  dominance.  The  great  portion  of  the 
population  was  grouped  in  the  lower  one  third  of  the  curve.  This  phenomenon 
suggests  the  existence  of  partial  dominance  of  the  "low  modifiers. " 

3  Selection  exercised  on  regions  1,2  and  3 in  the  sex 

chromosome  does  not  affect  the  crossover  value  of  the  region  outside  of  this, 
as  far  ae  we  can  see  from  our  data. 


KPOK 


*¥.'  ■ awr'TV 


:s^-t  ‘’^ir 

^ • . •.••■•>•  *»;■!■»■■  ...  • «T  -111  . ' w ' ■i'iikii;c*"ji»'  • S'.'"'  * '»  .»  . 'ledll 


. ^^' 


l(l  4wffl 


.tt*»fr.M\,;,  »•••»•'  «•««  ■ «w*  ■*■*>  ■'"■■■•  ’ ■ • "■  '^-v  ■ ■'^<r;.’.T  *■  ’ . m :■■  r ■ '■  ••  51 

•V  ^i.  • . ■ .;-•.  V ,,  Jf 

f Ip 


' .’^v , ,..£#■ 

y : L,V-S....,.,  ' ' ; ,%  ,w-.# ■■  ’■■'*■'■ 


\*Y.  .»  • 

; .%■ 
V'.®: 


~\  ii  ."V"  , > , ” s^.'  ..  . J ' i inr*  ^. 

■ ■■  '-•’♦•■».;■  . ^‘  ■ ■ Vii'A'.ftKr*..*^  ' ' .'■•  '..  o 


'?ie^ 


, , 1*.  .(«>).  •’1.*'' 


' f .,  ! H’t  ''.  V ' ^"***  ^*Eba  |p'  ^ 

*'  ■ I ' '‘  , V' ',  ' '■  i ' ■ •■  ( I ^ ''  ^ *'  ' ' 1 ■ /'  T 


(■'  , . i ' ' 1 ^1..  i.  ...  r'  » 


^ :».  ..  r' 


~ * ' • • \ ' ■ '■  . . • ■ ■•■•>..■“  Vi' t:^-M;}  ®'  ■•-.■'I 

^wa6^»^7 '<^■''**'411!^!^'?^ 

■ 


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7 <.'('! 


\ , ■ '.i. . ,' 


X^fK 

’■  ..I  . . 


I.  4, 


::i 


'St>'T\ 


f< 


X 


iJ^' 


'v  f,  . 

‘J 


,1..^ 


>■■■  rr^i, 

...W  ^ 


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■t 


...» 


,'Vfi 


^■.  - 

y Mi. 

■ ■w 


tfl  ' 


% 


,•»'.  i 


'jv 

u ^ - jjj-. 


.ll^  .. 


-17- 


V 

ACK'NDWLEDGEdENT . 

The  writer  is  very  grateful  to  Prof.  J.  A.  Detlefsen  for  the 
opportvinity  to  work  on  this  prohlera.  It  ia  with  a deep  sense  of  appreciation 
to  say  that  hut  for  his  helpful  suggestions  and  his  generosity  in  vc^ulping 
the  writer  with  necessary  laboratory  conveniences  this  work  would  not  have 
been  possible. 


-18- 


VI 


LITEUfiTUHE  CITED. 

Bridges,  C.  B.  1915  A linkage  variation  in  Drosophila.  Jour.  Expt.  Zool.  , 
vol.  19,  pp.  1-21 

Detlefsen,  J.  A.  and  Robert s £.  1921  Studies  on  Crossing  Over.  Jour.  £3cp. 

Zool.  vol.  32,  no.  2,  pp.  333-354. 


Detlefsen,  J.  A.  1920  Is  crossing  over  a function  of  distance?  Proc.  Nat'l. 
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